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The zona pellucida (plural zonae pellucidae, also egg coat or pellucid zone) is a specialized
extracellular matrix In biology, the extracellular matrix (ECM), also called intercellular matrix, is a three-dimensional network consisting of extracellular macromolecules and minerals, such as collagen, enzymes, glycoproteins and hydroxyapatite that provide s ...
that surrounds the
plasma membrane The cell membrane (also known as the plasma membrane (PM) or cytoplasmic membrane, and historically referred to as the plasmalemma) is a biological membrane that separates and protects the interior of all cells from the outside environment (t ...
of
mammal Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur ...
ian
oocyte An oocyte (, ), oöcyte, or ovocyte is a female gametocyte or germ cell involved in reproduction. In other words, it is an immature ovum, or egg cell. An oocyte is produced in a female fetus in the ovary during female gametogenesis. The female ...
s. It is a vital constitutive part of the oocyte. The zona pellucida first appears in unilaminar primary oocytes. It is secreted by both the oocyte and the
ovarian follicle An ovarian follicle is a roughly spheroid cellular aggregation set found in the ovaries. It secretes hormones that influence stages of the menstrual cycle. At the time of puberty, women have approximately 200,000 to 300,000 follicles, each with ...
s. The zona pellucida is surrounded by the corona radiata. The corona is composed of cells that care for the egg when it is emitted from the ovary. This structure binds
spermatozoa A spermatozoon (; also spelled spermatozoön; ; ) is a motile sperm cell, or moving form of the haploid cell that is the male gamete. A spermatozoon joins an ovum to form a zygote. (A zygote is a single cell, with a complete set of chromos ...
, and is required to initiate the acrosome reaction. In the mouse (the best characterised mammalian system), the zona glycoprotein,
ZP3 Zona pellucida sperm-binding protein 3, also known as zona pellucida glycoprotein 3 (Zp-3) or the sperm receptor, is a ZP module-containing protein that in humans is encoded by the ''ZP3'' gene. ZP3 is the receptor in the zona pellucida which bi ...
, is responsible for sperm binding, adhering to
protein Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, res ...
s on the sperm plasma membrane. ZP3 is then involved in the induction of the acrosome reaction, whereby a spermatozoon releases the contents of the acrosomal vesicle. The exact characterisation of what occurs in other species has become more complicated as further zona proteins have been identified. In humans, five days after the fertilization, the
blastocyst The blastocyst is a structure formed in the early embryonic development of mammals. It possesses an inner cell mass (ICM) also known as the ''embryoblast'' which subsequently forms the embryo, and an outer layer of trophoblast cells called th ...
performs zona hatching; the zona pellucida degenerates and decomposes, to be replaced by the underlying layer of
trophoblast The trophoblast (from Greek : to feed; and : germinator) is the outer layer of cells of the blastocyst. Trophoblasts are present four days after fertilization in humans. They provide nutrients to the embryo and develop into a large part of the p ...
ic cells. The zona pellucida is essential for oocyte growth and
fertilization Fertilisation or fertilization (see spelling differences), also known as generative fertilisation, syngamy and impregnation, is the fusion of gametes to give rise to a new individual organism or offspring and initiate its development. Pro ...
.


Structure

The zona pellucida is a translucent matrix of cross-linked
glycoprotein Glycoproteins are proteins which contain oligosaccharide chains covalently attached to amino acid side-chains. The carbohydrate is attached to the protein in a cotranslational or posttranslational modification. This process is known as glyco ...
filaments that surrounds the mammalian
oocyte An oocyte (, ), oöcyte, or ovocyte is a female gametocyte or germ cell involved in reproduction. In other words, it is an immature ovum, or egg cell. An oocyte is produced in a female fetus in the ovary during female gametogenesis. The female ...
and is 6.5–20 μm thick depending on the species. Its formation, which depends on a conserved Zona pellucida-like (ZP) module that mediates the polymerization of egg coat components, is critical to successful fertilization. In non-mammals it is called the vitelline membrane or vitelline envelope.


Function

The thick membrane of the zona pellucida functions to only allow species-specific fertilization; to prevent polyspermy, and enable the acrosome reaction for the successful adhesion and penetration by the
sperm cell Sperm is the male reproductive cell, or gamete, in anisogamous forms of sexual reproduction (forms in which there is a larger, female reproductive cell and a smaller, male one). Animals produce motile sperm with a tail known as a flagellum, ...
. Also allows correct
embryo development An embryo is an initial stage of development of a multicellular organism. In organisms that reproduce sexually, embryonic development is the part of the life cycle that begins just after fertilization of the female egg cell by the male sperm c ...
and size. The major glycoproteins of the egg coat responsible, are known as sperm-binding proteins. The four major sperm-binding proteins, or sperm-receptors, are ZP1, ZP2,
ZP3 Zona pellucida sperm-binding protein 3, also known as zona pellucida glycoprotein 3 (Zp-3) or the sperm receptor, is a ZP module-containing protein that in humans is encoded by the ''ZP3'' gene. ZP3 is the receptor in the zona pellucida which bi ...
, and ZP4. They bind to capacitated spermatozoa and induce the acrosome reaction. Successful
fertilization Fertilisation or fertilization (see spelling differences), also known as generative fertilisation, syngamy and impregnation, is the fusion of gametes to give rise to a new individual organism or offspring and initiate its development. Pro ...
depends on the ability of sperm to penetrate the
extracellular matrix In biology, the extracellular matrix (ECM), also called intercellular matrix, is a three-dimensional network consisting of extracellular macromolecules and minerals, such as collagen, enzymes, glycoproteins and hydroxyapatite that provide s ...
of the zona pellucida that surrounds the egg. In the mouse: *
ZP3 Zona pellucida sperm-binding protein 3, also known as zona pellucida glycoprotein 3 (Zp-3) or the sperm receptor, is a ZP module-containing protein that in humans is encoded by the ''ZP3'' gene. ZP3 is the receptor in the zona pellucida which bi ...
allows species-specific sperm binding * ZP2 mediates subsequent sperm binding * ZP1 cross-links ZP2 and ZP3. Data with native human protein are not currently available.


Immunocontraception

ZP module-containing glycoproteins ZP1, ZP2,
ZP3 Zona pellucida sperm-binding protein 3, also known as zona pellucida glycoprotein 3 (Zp-3) or the sperm receptor, is a ZP module-containing protein that in humans is encoded by the ''ZP3'' gene. ZP3 is the receptor in the zona pellucida which bi ...
and ZP4 are targets for immunocontraception in mammals. In non-mammals, the zona pellucida is called the vitelline membrane or envelope, and the vitelline envelope in
insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three pa ...
s, and plays an important role in preventing cross-breeding of different species, especially in species such as
fish Fish are Aquatic animal, aquatic, craniate, gill-bearing animals that lack Limb (anatomy), limbs with Digit (anatomy), digits. Included in this definition are the living hagfish, lampreys, and Chondrichthyes, cartilaginous and bony fish as we ...
that fertilize outside of the body. The zona pellucida is commonly used to control wildlife population problems by immunocontraception. When the zona pellucida of one animal species is injected into the bloodstream of another, it results in sterility of the second species due to
immune response An immune response is a reaction which occurs within an organism for the purpose of defending against foreign invaders. These invaders include a wide variety of different microorganisms including viruses, bacteria, parasites, and fungi which coul ...
. This effect can be temporary or permanent, depending on the method used. In
New Jersey New Jersey is a state in the Mid-Atlantic and Northeastern regions of the United States. It is bordered on the north and east by the state of New York; on the east, southeast, and south by the Atlantic Ocean; on the west by the Delawa ...
, immunocontraception using porcine zona pellucida has been trialled for the control of
deer Deer or true deer are hoofed ruminant mammals forming the family Cervidae. The two main groups of deer are the Cervinae, including the muntjac, the elk (wapiti), the red deer, and the fallow deer; and the Capreolinae, including the re ...
.


Additional images

File:Gray9.png, First stages of segmentation of a mammalian
ovum The egg cell, or ovum (plural ova), is the female reproductive cell, or gamete, in most anisogamous organisms (organisms that reproduce sexually with a larger, female gamete and a smaller, male one). The term is used when the female gamete i ...
File:Gray1164.png, Section of vesicular ovarian follicle of cat, x 50 File:HumanEmbryogenesis.svg, The initial stages of
human embryogenesis Human embryonic development, or human embryogenesis, is the development and formation of the human embryo. It is characterised by the processes of cell division and cellular differentiation of the embryo that occurs during the early stages of de ...


References


Further reading

* * * * Bagnell C (2005). "Animal Reproduction". Rutgers University Department of Animal Sciences. * * * * * *


External links

* - "Female Reproductive System: ovary, cumulus oophorus " * - "Female Reproductive System: ovary, multilaminar primary follicle" * - "Mammal, canine ovary (LM, High)"
Image at um.edu.mt

Image at um.edu.mt
{{DEFAULTSORT:Zona Pellucida Germ cell structures Reproductive system Vertebrate developmental biology